Ocular pigmentation in humans, great apes, and gibbons is not suggestive of communicative functions

Pigmentation patterns of the visible part of the eyeball, encompassing the iris and portions of the sclera, have been discussed to be linked to social cognition in primates. The cooperative eye hypothesis suggests the white sclera of humans to be a derived adaptive trait that enhances eye-mediated communication. Here, we provide a comparative analysis of ocular pigmentation patterns in 15 species of hominoids (humans, great apes & gibbons) that show marked differences in social cognition and quantify scleral exposure at the genus level. Our data reveals a continuum of eye pigmentation traits in hominoids which does not align with the complexity of gaze-mediated communication in the studied taxa. Gibbons display darker eyes than great apes and expose less sclera. Iridoscleral contrasts in orangutans and gorillas approach the human condition but differ between congeneric species. Contrary to recent discussions, we found chimpanzee eyes to exhibit a cryptic coloration scheme that resembles gibbons more than other apes. We reevaluate the evidence for links between social cognition and eye pigmentation in primates, concluding that the cooperative eye hypothesis cannot explain the patterns observed. Differences in scleral pigmentation between great apes and humans are gradual and might have arisen via genetic drift and sexual selection.

Ocular pigmentation in humans, great apes, and gibbons is not suggestive of communicative functions - having an eye on the ‘cooperative eye hypothesis’

Pigmentation patterns of the visible part of the eyeball, encompassing the iris and portions of the sclera, have been discussed to be linked to social cognition in primates. In the context of the cooperative eye hypothesis, the white sclera of humans has been viewed as a derived adaptive trait, enhancing communication via glance cueing. Here, we provide a comparative analysis of ocular pigmentation patterns in 15 species of hominoids (humans, great apes & gibbons) representing all extant ape genera, based on photographs and literature data. Additionally, we quantify hominoid scleral exposure on the genus level during different glancing situations. Our data reveals a continuum of eye pigmentation traits among the studied taxa. Gibbons display darker, more uniformly colored eyes than great apes and expose less sclera, particularly during averted glancing. Iridoscleral contrasts in orangutans and gorillas approach the human condition but differ between congeneric species. Contrary to recent discussions, we found chimpanzee eyes to exhibit a cryptic coloration scheme that resembles gibbons more than other great apes and that does not enhance glance cueing or gaze following. We critically evaluate the evidence for links between social cognition and eye pigmentation in primates, concluding that the cooperative eye hypothesis cannot convincingly explain the patterns observed. Although the human eye exhibits unique traits that are likely linked to social communication, high iridoscleral contrast is not one of them. Differences in scleral pigmentation between great apes and humans are gradual and might have arisen via genetic drift and sexual selection.

A new Norwegian Lemming subspecies from Novaya Zemlya, Arctic Russia

Norwegian Lemming Lemmus lemmus is a remarkable population cycling species having a number of aposematic traits in coloration and behavior. This species was thought to be the only mammal endemic to Fennoscandia. Here, we report on the discovery of a distinct lineage of this species from Novaya Zemlya. This lineage is described here as the new subspecies Lemmus lemmus chernovi ssp. nov. that morphologically differs from the nominate subspecies by having a cryptic coloration. Our time-calibrated phylogeny revealed that this insular subspecies was isolated there since the Eemian interglacial (mean age 93 Ka). Hence, bright coloration and antipredator behavior of the nominate subspecies are novel aposematic traits that evolved since its isolation in a European refugium. This new discovery indicates that Novaya Zemlya supported a cryptic polar refugium for cold-adapted terrestrial fauna during the Pleistocene. Finally, our findings highlight that allopatric ranges of the true lemmings (Lemmus) reflect stepwise range expansions in cold climatic episodes with subsequent isolation in refugia during interglacial events.

Biased movement drives local cryptic coloration on distinct urban pavements

Explanations of how organisms might adapt to urban environments have mostly focused on divergent natural selection and adaptive plasticity. However, differential habitat choice has been suggested as an alternative. Here, we test for habitat choice in enhancing crypsis in ground-perching grasshoppers colonizing an urbanized environment, composed of a mosaic of four distinctly coloured substrates (asphalt roads and adjacent pavements). Additionally, we determine its relative importance compared to present-day natural selection and phenotypic plasticity. We found that grasshoppers are very mobile, but nevertheless approximately match the colour of their local substrate. By manipulating grasshopper colour, we confirm that grasshoppers increase the usage of those urban substrates that resemble their own colours. This selective movement actively improves crypsis. Colour divergence between grasshoppers on different substrates is not or hardly owing to present-day natural selection, because observed mortality rates are too low to counteract random substrate use. Additional experiments also show negligible contributions from plasticity in colour. Our results confirm that matching habitat choice can be an important driver of adaptation to urban environments. In general, studies should more fully incorporate that individuals are not only selective targets (i.e. selected on by the environment), but also selective agents (i.e. selecting their own environments).

Antipredator strategies of pupae: how to avoid predation in an immobile life stage?

Antipredator strategies of the pupal stage in insects have received little attention in comparison to larval or adult stages. This is despite the fact that predation risk can be high during the pupal stage, making it a critical stage for subsequent fitness. The immobile pupae are not, however, defenceless; a wide range of antipredator strategies have evolved against invertebrate and vertebrate predators. The most common strategy seems to be ‘avoiding encounters with predators' by actively hiding in vegetation and soil or via cryptic coloration and masquerade. Pupae have also evolved behavioural and secondary defences such as defensive toxins, physical defences or deimatic movements and sounds. Interestingly, warning coloration used to advertise unprofitability has evolved very rarely, even though the pupal stage often contains defensive toxins in chemically defended species. In some species, pupae gain protection from conspecifics or mimic chemical and auditory signals and thereby manipulate other species to protect them. Our literature survey highlights the importance of studying selection pressures across an individual's life stages to predict how ontogenetic variation in selective environments shapes individual fitness and population dynamics in insects. Finally, we also suggest interesting avenues for future research to pursue. This article is part of the theme issue ‘The evolution of complete metamorphosis’.

Evaluation of the effectiveness of cryptic coloration of the Carolina anole’s skin

Cryptic coloration of animals’ integument is one of the effective adaptations that allow them to lead an active lifestyle while being protected from natural enemies due to visual disguise. This is achieved by the similarity of body color of a particular individual to the background of various substrates in its environment. The morphological and functional basis of cryptic coloration in vertebrates, including reptiles, is ensured by the skin pigmentation. Using bioinformatic methods, we calculated the skin camouflage index of the Carolina anole (Anolis carolinensis Voigt, 1832) in various conditions of its habitat. The skin camouflage index (Ic) is the ratio of the sum of the average values of rgb coordinates of the skin color to the sum of the average values of rgb coordinates of the color of the external substrate. Ic satisfies the effective level of adaptation to habitat conditions if it falls within the range of 0.80–1.20. It has been shown that rgb-values of the dominant color of the dorsal skin of green anoles slightly differ from the similar parameters characteristic of the deciduous habitat, which is reflected by Ic of its skin (0.94). In the brown anoles on a background of woody substrate, the Ic value of its skin (0.88) is also optimal, since it exceeds the lower limit (0.80), which indicates close values of the sums of the rgb coordinates of the skin color and the color of the external background. In the mixed green-brown anoles, the deciduous habitat is preferable to the woody one. In the first case, Ic (1.11) is in the optimum zone, and in the second case, Ic (0.70) goes beyond the lower limit of the optimum, which indicates a greater vulnerability of the animal to external threats. We have confirmed the relativity of visual hiding of the skin that is effective only in the habitat conditions in which the camouflage abilities of the skin manifest as fully as possible.

Flight initiation distance, color and camouflage

Camouflage is widespread throughout the animal kingdom allowing individuals to avoid detection and hence save time and energy rather than escape from an approaching predator. Thus, camouflage is likely to have co-evolved with antipredator behavior. Here, we propose that camouflage results in dichotomous escape behavior within and among species with classes of individuals and species with cryptic coloration having shorter flight initiation distances (FIDs; the distance at which an individual takes flight when approached by a human). We report the results of 2 tests of this hypothesis. First, bird species with cryptically colored plumage have consistently shorter FID than closely related species without such color. Within species with sexually dimorphic plumage, brightly colored adult male common pheasants Phasianus colchicus and golden pheasants Chrysolophus pictus have long and variable FID, whereas cryptically colored juveniles and adult females have short and invariable FID. Second, FID in females was predicted by presence or absence of cryptic color, FID in males and their interaction. These findings are consistent with the hypothesis that risk-taking behavior has been attuned to camouflage, and that species with different levels of camouflage differ consistently in their FID.

Morphology of the immature stages and notes on the natural history of the little-known plume moth Stenoptilodes juanfernandicus Gielis (Lepidoptera: Pterophoridae)

Stenoptilodes juanfernandicus Gielis, 1991 (Lepidoptera: Pterophoridae) was described from the Juan Fernandez Islands, Chile; it was subsequently recorded from the Galápagos Islands and mainland Ecuador. Its larva and pupa are described and illustrated for the first time with the aid of light and scanning electron microscopy. Descriptions are based on specimens collected as larvae displaying cryptic coloration feeding on inflorescences of Verbena hispida Ruiz & Pav. (Verbenaceae) in the Andes of northern Chile. Several differences were found with the larva and pupa of congeneric representatives, suggesting that immature morphology should be further explored to assess its value in the taxonomy and systematics of Stenoptilodes Zimmerman, 1958. Verbena hispida is the first host plant recorded for S. juanfernandicus, a finding that will be helpful to further explore different Neotropical environments to better characterize the geographic range of this plume moth.